You are on page 1of 5

HORTSCIENCE 52(8):1060–1064. 2017. doi: 10.

21273/HORTSCI12058-17 have an alternated arrangement forming


a spiriform sequence. The plants have long
Systems of Pruning on Jigacho pendular floral whorls, with female, male,
and hermaphrodite flowers, differenti-
(Vasconcellea stipulata Badillo) under ated by the androecium and gynoecium
(Kubitzki, 2003). Flowers are yellow with
reddish areas (Scheldeman et al., 2011).
Greenhouse Conditions Fruits are small, 8–10-cm long and 4–5-cm
wide, yellow, weighing between 40 and
Borja Velazquez-Martí 1 150 g. They have five lobes and grooves
Departamento de Ingeniería Rural y Agroalimentaria, Universitat (6–10) that give a star shaped cross sec-
Polit
ecnica de Val
encia, Camino de Vera s/n, 46022 Valencia, Spain tion, being pentacarpelar. The skin of the
fruit is usually soft and easy to peel, its
Marco Castillo and Fidel Rodríguez flesh is creamy, yellow, and smooth, with
Facultad de Ciencias Agropecuarias, Universidad T ecnica de Ambato, seven or 10 seeds.
Vasconcellea ssp. are trees grown not
Campus Querochaca, canton Cevallos, Tungurahua 18-01-334, Ecuador only for fruit consumption, but also as
Isabel L
opez-Cortes ornamental plants noted for the exquisite floral
fragrance. Vasconcellea stipulate Badillo
Departamento de Produccion Vegetal, Universitat Polit
ecnica de Val
encia, has been traditionally cultivated in domestic
Camino de Vera s/n, 46022 Valencia, Spain form on small plots, and its commercial
development has been limited (Badillo,
Stefany Alcivar-Bastidas 1993; Scheldeman et al., 2002; Van den
Facultad de Ingeniería, Universidad Cat
olica de Santiago de Guayaquil, Av. Eynden et al., 2003). Nevertheless, its good
Carlos Julio Arosemena km. 1 1/2 Vía Daule, Guayaquil, Ecuador organoleptic qualities give it a high potential
in the markets of Europe and North America
Additional index words. Ecuador, Andes, toronche, Vasconcellea stipulata, underutilized where it is considered an exotic fruit (Baeza
fruits species, crop cycle, crop management system et al., 1990; Dhuique Mayer et al., 2001;
Abstract. Vasconcellea stipulata Badillo is a fruit tree native of the temperate areas of the Scheldeman et al., 2002). The fruit can be
Andes valleys, found between 2000 and 2600 m above sea level. The good organoleptic eaten fresh or processed industrially into
qualities and few seeds of its fruit, compared with those of other Caricas, give it a high jellies, jams, and juices, usually mixed with
potential in markets. The development of this crop would allow the economic development other fruits (Cossio, 1988). Its fruit is also
of small-scale farmers and contribute to the preservation of genetic resources native to the interesting because of its high papain con-
Andean region. However, there are few studies concerning its appropriate cultivation. The tent (Kyndt et al., 2007; Misas-Villamil
aim of this study was to analyze the effect of different pruning techniques under greenhouse et al., 2016; Scheldeman et al., 2003), which
conditions. Trees were pruned to one, two, or three axes, or not pruned (control). The is used in the pharmaceutical industry for the
experiment was performed in a tunnel greenhouse with polyethylene cover, located 2950 m treatment of cancerous tumors and lymph,
above sea level in Querochaca (Ecuador), with an annual rainfall of 465 mm and an average herniated discs, and abnormal formations
annual temperature of 12.9 8C. The duration of each stage of development was recorded. that occur in the arteries (atherosclerosis).
The number of flower clusters, number of flowers, production, incidence of diseases, and Papain is an anti-inflammatory agent in
weight and size of fruit were recorded. Cuttings took over 90 days for getting before being cases of infections and trauma (Aravind
transplanted into the greenhouse. Flowering began 283 days (40 weeks) after trans- et al., 2013; Flindt, 1978; Kyndt et al.,
plantation; fruit set began at 30 days after flowering (44 weeks after transplant); and the 2007). It is also used for healing ulcers in
fruit started to mature 170 days after fruit set (69 weeks after transplant). The plants the digestive tract and in the treatment of
pruned to three axes had the greatest number of flowers and fruits and the greatest dyspepsia and kidney dysfunction. It has
production, with an average 54 fruits per tree. Pruning had no effect on fruit dimensions. industrial and cosmetic use in healing and
eczema correction. In food industry, it is
used as a meat tenderizers (Endt, 1983;
Jigacho or toronche (Vasconcellea stipu- scars on the stem (Kubitzki, 2003). Leaves Cossio, 1988).
lata Badillo) is a fruit native to temperate are polylobed, pentagon-webbed, with The development of this crop would
areas of the Andes valleys, found between a very high polymorphism (Fig. 1). They allow the economic development of small
2000 and 2600 m above sea level. This
species is cultivated mainly in southern
Ecuador and northern Peru (Badillo, 1993;
Scheldeman et al., 2007). It is a semiwoody
tree, which reaches 5–10 m height. It is
characterized by spines (spiniform stipules)
in the stem and leaf petiole. When leaves
abscise, the petiole insertion produces large

Received for publication 27 Apr. 2017. Accepted


for publication 22 May 2017.
The participation of Borja Velazquez Martí in
this work was possible by the funding from the
Ecuadorian Government through PROMETEO
program, leaded by the Secretaría Nacional
de Educaci on Superior, Ciencia y Tecnología
(SENESCYT).
1
Corresponding author. E-mail: borvemar@dmta. Fig. 1. Morphology of Jigacho (Vasconcellea stipulata V.M. Badillo) fruit and leaf. (A) Fruit;
upv.es. (B) transversal section of the fruit; (C) seed (D) abaxial connection of leaf-petiole; (E) leaf.

1060 HORTSCIENCE VOL. 52(8) AUGUST 2017


farmers and contribute to the preservation 11 d under shade, time in which part of the sequently, a continuous pruning was per-
of genetic resources originated in the latex was eliminated and the cuts healed. formed to remove unwanted shoots until the
Andean region. Its cultivation under green- After this time, the cuttings were placed on plant reached a year after sprouting.
house condition at high altitude is an rooting medium for 3 months. The substrate Fertilizers were applied before planting
opportunity to increase production under was composed of 40% topsoil, 40% humus, with 100 g·m–2 triple superphosphate and 100 g
controlled conditions. However, there are 20% pomina. The cuttings were then planted muriate of potash/m2. After planting, 26 g
few studies concerning proper cultural in a greenhouse tunnel under polyethylene N, 50 g P2O5 and 33 g K2O/plant were
management. The aim of this study was to cover in sandy loam soil, irrigated by a drip applied after 4, 7, and 10 months. Tempera-
analyze the effect of different pruning system (Fig. 2). The greenhouse was located tures in the greenhouse average 21.0 C, with
techniques in the development of jigacho at the Experimental Farm of the Technical 9.6 C minimum and 33.4 C maximum. The
cultivated in a greenhouse. We studied their University of Ambato (Ecuador), located at average relative humidity in the greenhouse
vegetative development and influence of 116#27.31$, 7834#32.42$, X = –1,369,846 m, was 57%, with 26% minimum and 94%
pruning on the number of flowers, produc- Y = –78,607,242 m, at 2950 m above sea maximum.
tion, weight and size of fruits, as well as level. According to Querochaca Meteorolog- To evaluate the effect of pruning, the
susceptibility to the most common diseases ical Station (2011–2014), the climate in this following parameters were measured:
of this species. area is temperate, with 12 C minimum
temperature, 18 C maximum temperature.  Number of days to first flower
Materials and Methods The average temperature of 12.9 C; 250 mm  Number of days to start fruiting
minimum and 600 mm maximum precipita-  Number of days to start harvest
Plant material. For the study, cuttings of tion, the average annual precipitation was  Number of fruits per plant
jigacho stem (Vasconcellea stipulata Bad- 465 mm; relative humidity of 77%; and  Number of fruits per cluster
illo) (Fig. 1) were taken from homogeneous a speed of wind of 4.2 m/s.  Polar diameter of the fruit
adult plants 5 years old, located in the Experimental design. The experiment was  Equatorial diameter of fruit
cantons Salcedo and Cevallos (Ecuador). conducted over 4 years. The most important  Soluble solids (Brix)
The length of the cuttings was 25–30 cm approach to pruning jigacho is to define tree  Average fruit weight
(3–4 internodes). The cuttings were kept for shape. This pruning approach determines the  Observation of the incidence of pests and
number of branches and its structure. After diseases.
establishment, no significant changes are
made in the structure of the plant. The effect Comparison of pruning methods was per-
of pruning was studied in this work, which formed using analysis of variance (ANOVA)
can influence production. Four different with a confidence level of 95%.
types of pruning were studied: S1, the plant
grew forming only one axis; S2, the plants Results and Discussion
were pruned forming two axes without
branching; S3, the plants were pruned in Analysis of the growth cycle. Figure 4
Fig. 2. View of jigacho planted in greenhouse: S1
three axes without branching; C, the plants shows the variation in the average number
(one axis) and S2 (two axes). had free growth. These treatments were of flower clusters in the trees with the
randomly distributed in five blocks. Thus, different types of pruning during the weeks
20 experimental plots were analyzed each after transplant. Flowering began at 283 d
year. In each block, a different type of after planting. The curves representing the
pruning was performed in a plot. The distri- variation of number of flower clusters vs.
bution of the plot in the greenhouse is shown time were sigmoid. Flowering was initially
in Fig. 3. Each plot was composed of 12 slow; then, it increased exponentially be-
plants. The plants were separated 1.5 m each tween weeks 40 and 55. The coefficients of
on a row and the rows were 1.4 m distanced; determination of the exponential curves in
therefore, each block occupied 25.2 m2. those weeks vary between 0.97 and 0.98 for
The plants remained in the greenhouse for S1 and S2 treatment. The worse exponen-
4 years. tial curve represents the flowering process
Fig. 3. Distribution of sampled plots in the On each block, pruning was performed vs. time in the S3 and control treatment,
greenhouse. 3 months after the stakes were planted. Sub- which have no differences. These treat-
ments produce a more lineal flowering pro-
cess. Subsequently, increasing the bloom
decreases until the number of flower clusters
is constant. This model allows analyzing
differences between treatments and predict-
ing the production with few weeks of the
cycle because the number of flowers and fruit
is proportional. These equations are not lineal
probably because of liberation of the hor-
mones to flowering are progressive. Table 1
shows the means and standard deviations of
the number of clusters at the end of each
month. Analysis of variance was performed
to determine if there are differences between
the numbers of floral clusters in different
types of pruning with 95% of confidence
level, the number of flowers per plant with
time, number of fruits per tree with time,
number of drop fruits over time.
The number of flower clusters per plant
Fig. 4. Variation in the number of floral clusters in different types of pruning. formed at 283 and 313 d showed narrow

HORTSCIENCE VOL. 52(8) AUGUST 2017 1061


Table 1. Average and SD of evaluated parameters according to the time after transplantation (sample 48 plants, 12 per year). Treatments—S1: plants were growing
forming only one axis; S2: plants were pruned forming two axes without branching; S3: plants were pruned in three axes without branching; C: plants had free
growth. Analysis of variance at 95% confidence level.
Week 40 Week 44 Week 48 Week 52 Week 56 Week 60 Week 64 Week 68
283 d 313 d 343 d 373 d 403 d 433 d 463 d 493 d
Number of floral clusters per plant
S1 6.11 ± 2.30 a 12.79 ± 3.33 a 22.19 ± 4.85 a 32.83 ± 5.52 a 43.30 ± 6.78 a 52.90 ± 9.02 a 59.50 ± 19.27 a 62.35 ± 15.28 a
S2 6.13 ± 4.45 a 13.08 ± 8.04 a 25.18 ± 8.37 a 38.45 ± 7.61 a 52.20 ± 8.83 a 60.95 ± 5.66 a 66.65 ± 12.18 a 66.20 ± 16.47 a
S3 3.39 ± 2.00 b 7.36 ± 3.35 b 12.17 ± 4.48 b 17.78 ± 6.26 b 23.55 ± 7.43 b 29.60 ± 8.76 b 34.05 ± 6.68 b 35.75 ± 14.99 b
C 2.29 ± 2.83 b 6.57 ± 5.86 b 12.04 ± 8.49 b 18.70 ± 11.38 b 25.85 ± 14.25 b 32.45 ± 17.18 b 35.73 ± 11.49 b 36.40 ± 11.28 b
Number of flowers per plant
S1 6.76 ± 2.54 a 14.11 ± 5.88 a 30.96 ± 10.04 a 31.63 ± 9.81 a 42.10 ± 9.19 b 50.50 ± 12.47 a 44.50 ± 9.81 a 33.95 ± 10.16 a
S2 6.96 ± 5.12 a 13.62 ± 8.88 a 31.65 ± 15.14 a 38.53 ± 11.46 a 58.42 ± 9.97 a 66.55 ± 8.97 a 56.3 ± 10.79 a 36.95 ± 8.85 a
S3 4.35 ± 3.04 a 9.07 ± 5.15 a 17.83 ± 9.59 ab 17.95 ± 9.64 b 24.64 ± 8.90 c 26.35 ± 8.94 c 28.4 ± 10.04 b 22.70 ± 13.40 b
C 2.55 ± 2.93 a 6.47 ± 5.96 a 13.46 ± 9.42 b 14.82 ± 8.07 b 26.20 ± 12.75 c 30.05 ± 11.43 c 28.25 ± 12.00 b 18.72 ± 7.71 b
Number of fruits per plant
S1 0.00 ±.0.00 a 1.61 ± 1.03 a 5.19 ± 2.79 a 23.33 ± 9.81 a 36.25 ± 9.10 a 44.50 ± 12.45 a 52.6 ± 17.75 a 55.20 ± 23.05 a
S2 0.19 ± 0.42 a 1.64 ± 1.75 a 4.99 ± 3.01 a 24.13 ± 11.46 a 33.93 ± 11.15 a 40.20 ± 7.64 a 45.2 ± 7.56 a 49.45 ± 7.13 a
S3 0.05 ± 0.11 a 1.02 ± 0.89 a 4.39 ± 1.94 a 17.52 ± 9.64 a 25.40 ± 12.32 a 33.00 ± 17.94 a 38.35 ± 21.80 a 40.50 ± 8.42 b
C 0.00 ± 0.00 a 0.41 ± 0.52 a 3.04 ± 3.57 a 12.55 ± 10.52 a 17.08 ± 13.63 a 23.1 ± 7.09 b 27.90 ± 21.04 a 31.96 ± 10.74 b
Number of drop fruit
S1 — — — 0.83 ± 0.73 ab 1.08 ± 0.56 ab 7.65 ± 3.99 ab 3.60 ± 2.27 b 2.35 ± 0.80 b
S2 — — — 1.23 ± 0.32 a 1.45 ± 0.60 a 11.08 ± 4.64 a 6.70 ± 1.57 a 4.25 ± 1.31 a
S3 — — — 0.30 ± 0.29 b 0.53 ± 0.37 b 4.20 ± 2.21 b 1.50 ± 0.50 c 1.15 ± 0.57 b
C — — — 0.60 ± 0.35 ab 0.70 ± 0.47 b 7.25 ± 3.64 ab 2.89 ± 1.07 bc 2.16 ± 1.06 b

differences in the types of pruning applied.


This means that the four systems had small
different responses to the formation of flower
clusters. However, by the 48th week after
transplant, the plants pruned to S1 and S2
(principal axes 1 and 2) had statistically more
flowers than S3 treatment (main axis) and
control (free growth). The average number of
flowers in S1 and S2 treatments reached
64.5 per plant; however, in S3 and control
treatments, it reached only 36.3 flower
clusters per plant.
Figure 5 shows the variation in the
number of flowers per plant vs. time, and
in Table 1 is shown the ANOVA carried out
to evaluate the differences between the
treatments. Again it is shown that treatments
S1 and S2 have provided more flowers per Fig. 5. Variation in the number of flowers per plant with time (sample 60 plants, 12 per block).
plant. Of course, this is related with the
number of clusters. The greatest number of
flowers was obtained at 60th week. The trees
with more flowers are those who had un-
dergone the S2 treatment with an average of
66.55 flowers per plant. At the 54th and
64th week, the number of flowers decreased,
because of floral abscission and develop-
ment of fruit.
Figure 6 shows the number of fruit in the
trees for each type of pruning. It can be seen
that the number of fruits per tree grew
exponentially between the 44th and
55th week. From week 55, the number of
fruits decreased because of the increase in
fruit drop (Fig. 7). The types of pruning with
more fruits were S1 and S2, with 53 fruits
average at 69th week after transplantation.
These treatments differed significantly from
S3 and control treatments, which had a mean
of 38 fruits per tree, although pruning types Fig. 6. Variation of the number of fruits per tree with time (sample 60 plants, 12 per block).
S1 and S2 are also where there was more
fruit drop. pying a volume of 28.2 cm3, and average pruning. The size and weight in the fruit
The fruit reached the proper maturity for weight was 290 g, with a pH 4.49 and a juice grown in the greenhouse conditions were
harvest at 69th week after transplant. The with 9.79 Brix. No significant differences higher than those reported by Badillo (1993).
fruit were collected with average size of existed in the properties of the fruit be- According to the analysis presented, the
6.1 cm diameter and 16.5 cm long, occu- tween the different treatments of tested cycle followed the next pattern (Fig. 8): 90 d

1062 HORTSCIENCE VOL. 52(8) AUGUST 2017


Kumar et al. (2014), studied the effects of
tree spacing (5 · 2 m, 5 · 2.5 m, 5 · 3 m, 5 ·
3.5 m, and 5 · 4 m) and pruning (eight buds/
cane, six buds/cane, and four buds/cane) on
vegetative growth, physiological parameters,
fruit yield, and quality were studied in fig
(Ficus carica L.) cv. Deanna in the third and
fourth year of its growth during the period
2010–12. They observed that with increase in
tree spacing, growth parameters such as leaf
number, shoot length, internode length, tree-
spread, tree height, and tree circumference,
along with fruit yield both in terms of fruit
number and fruit weight per tree, declined
gradually under different pruning levels.
Increase in pruning level from eight buds/
cane to four buds/cane resulted in increased
leaf number, shoot length and internode
Fig. 7. Variation of the number of drop fruits over time (sample 60 plants, 12 per block). length. Yield characters, viz., fruit number/
tree, fruit weight/tree, fruit number/hectare,
and fruit weight/hectare were marginally
influenced by pruning. However, interaction
effects of pruning and spacing were found to
be nonsignificant. Effects of spacing were
more conspicuous than those of pruning. Best
results for maintenance of vigor and fruit
yield were observed under a spacing of 5 ·
2 m or 5 · 2.5 m, and four buds/cane pruning.
Although there was reduction in average fruit
Fig. 8. Cycle of Vasconcellea stipulata Badillo. size under closer spacing when compared
with wide spacing, fruit quality attributes
such as total soluble solids and acidity were
Table 2. Properties of the harvested fruits (sample At the 10th month after planting, the unaffected by various treatments. Effects of
400 fruits, 100 per year). presence of ‘‘brown leaf spot’’ was also closer spacing on growth and yield param-
x ± s detected. This was probably incited by fun- eters were more pronounced in the third
Equatorial diameter (cm) 6.1 ± 0.6 gus Asperisporium spp. This is a little para- year as compared with the fourth year,
Length (cm) 16.1 ± 1.2 site known by its specific attacks to the showing better response to treatments in
Volume (cm3) 282.2 ± 46.7 Caricaceae family, causing small necrotic young trees.
Weight (g) 289.5 ± 47.8 spots (0.2–0.8 cm in diameter) in the leaves’
Juice pH 4.51 ± 0.7 bundle. Under the leaf, there are also small
Soluble solids of the juice (Brix) 9.97 ± 0.42 Conclusions
necrotic spots randomly distributed.
Differences of disease affectation were not Cycle of Vasconcellea stipulata Badillo
observed between the studied pruning systems. cultivated in cold Andinean areas under
were spent for rooting and transplanting of Comparison with papaya. No previous greenhouse without additional heat system has
stem cuttings. After transplantation, the flow- publication has reported on the pruning of been described. It was found that flowering in
ering began at 283 d (40th week), fruit setting Vasconcellea spp., although they are very jigacho begins in at week 40 after transplan-
started at 30 d after flowering (44 weeks after proximate to Caricaceae. The most studied tation. The fructification begins at week 44
transplantation). The maturation time of the Carica species has been the pawpaw tree after transplanting. The maturation time of the
fruit from the fructification to harvest was (Carica papaya L.). Niklas and Thomas fruit is of 170 d, being harvested from week
170 d, performing the harvesting at 69 weeks (2007) studied the effects of its domestica- 69 after transplantation.
after transplantation. Table 2 shows mean tion on vegetative growth of this crop, but In this article, three types of pruning
and standard deviation of the parameters of this is a Caricaceae, cultivated in warm were assessed, comparing with the free
the harvested fruits. weather. Barros et al. (2009) studied the growth. It has been shown that plant con-
Diseases observed in the crop. A small effect of pruning at different heights of the figurations in one and two axes provide
infection of root rot was observed, which was trunk in the papaya cv. Golden, with two and a higher production than free development
attributed to the presence of Fusarium spp. It half years old, during the productive period. or the growing with three axes. This may be
was found at the level of the crown that the The treatments consisted of pruning at five because nutrients are better distributed and
color of the tissues turned lightly brown, and heights (0.5, 1.0, 1.5, 2.0 m starting from the differentiation hormone act more effi-
as the disease progressed, it became watery. ground, and complete without the apex). ciently. When the plant has free growing
The leaves turned chlorotic, being withered Evaluated characteristics were number of and the number of branches is high, the flower
until they fall, and also the fruit, until finally shoots at 25 and 50 d after the pruning, number of available flowers is reduced, and
the entire plant died. length of the sprouts longer than 5 cm at 50 d excess flowers make them less viable.
Another significant infection was in- after the pruning, and distribution of the The fruits obtained in trees under green-
cited by the presence of mildew on the shoots in trunk height at 50 d. These works house are larger and heavier than those
underside of leaves with chlorotic spots in demonstrated that Caricaceae production is obtained outdoors. The different types of
the beam, which enlarge and grouped, the influenced by pruning, whereas the different pruning did not influence the size and weight
photosynthetic area of the plant being types of pruning were shown not to change of the fruit. The disease infections in the
notably reduced. This probably occurred the fruit quality. Such finding has been greenhouse cultivation are acceptable for
by the presence of oidium. The attack verified in the present study, where pruning large-scale production.
became more severe in low relative humid- S1 and S2 improved the fruit yield of This article demonstrates that cultiva-
ity (Vivas et al., 2016). jigacho. tion of jigacho can be carried out under

HORTSCIENCE VOL. 52(8) AUGUST 2017 1063


greenhouse with an acceptable yield of 38 Kubitzki, K. 2003. Caricaceae. Flowering potential of Andean fruit crops exploring their
fruits per tree, about 11 kg. plants·dicotyledons. The families and genera centre of origin. Proc. Intl. Symp. on Sustainable
of vascular plants. Vol. 5, Springer, p. 57–61. Use of Plant Biodiversity to Promote New
Kumar, R., S. Ganesh, R. Chithiraichelvan, K.K. Opportunities for Hort. Production Dev. Intl.
Literature Cited Upreti, and V.V. Sulladmath. 2014. Effect of Soc. Hort. Sci., The Hague, The Netherlands.
spacing and pruning on growth, yield and p. 97–102.
Aravind, G., D. Bhowmik, S. Duraivel, and G. Harish. quality of cv. Deanna fig (Ficus carica L.). J. Scheldeman, X., P. Van Damme, and J. Romero
2013. Health benefits and medicinal properties of Hort. Sci. 9(1):31–37. Motoche. 2002. Highland papayas in southern
Carica papaya. Ann. Plant Sci. 2(2):1–27. Kyndt, T., J.M. Van Damme Els, J. Van Beeumen, Ecuador: Need for conservation actions. Acta
Badillo, V.M. 1993. Caricaceae. Segundo esquema. and G. Gheysen. 2007. Purification and charac- Hort. 575(1):199–205.
Rev. Fac. Agron. Univ. Cent. Venez. 43:1–111. terization of the cysteine proteinases in the latex Scheldeman, X., L. Willemen, G. Coppens
Baeza, G., D. Correa, and C. Salas. 1990. Pro- of Vasconcellea spp. FEBS J. 274(2):301–574. d’Eeckenbrugge, E. Romeijn-Peeters, M.T.
teolytic enzymes in Carica candamarcensis. J. Misas-Villamil, J.C., R.A.L. van der Hoorn, and G. Restrepo, J. Romero Motoche, D. Jimenez, M.
Sci. Food Agr. 51:1–9. Doehlemann. 2016. Papain-like cysteine pro- Lobo, C.I. Medina, C. Reyes, D. Rodríguez, J.A.
Barros, F.L.D., E.R. Schmildt, J.A.T. do Amaral, teases as hubs in plant immunity. New Phytol. Ocampo, P. Van Damme, and P. Goetgebeur.
and R.I. Coelho. 2009. Influence of the pruning 212:902–907. 2007. Distribution, diversity and environmental
to different heights in papaya tree ‘Golden’. Niklas, K.J. and E.M. Thomas. 2007. Carica adaptation of highland papayas (Vasconcellea
Rev. Ci^enc. Agron. 40(4):596–601. papaya (Caricaceae): A case study into the spp.) in tropical and subtropical America. Bio-
Cossio, F. 1988. Il Babaco. Edagricoles, Bologna, effects of domestication on plant vegetative divers. Conserv. 16:1867–1884.
Italy. growth and reproduction. Amer. J. Bot. 94 Van den Eynden, V., E. Cueva, and O. Cabrera.
Dhuique Mayer, C., Y. Caro, M. Pina, J. Ruales, M. (6):999–1002. 2003. Wild foods from Ecuador. Econ. Bot. 57
Dornier, and J. Graille. 2001. Biocatalytic Scheldeman, X., T. Kyndt, G. Coppens (4):576–603.
properties of lipase in crude latex from babaco d’Eeckenbrugge, R. Ming, R. Drew, and Vivas, M., H.C.C. Ramos, P.H.D. Santos, S.F.
fruit (Carica pentagona). Biotechnol. Lett. 23 P. Van Damme. 2011. C. Kole (ed.). Wild Silveira, T.N.S. Pereira, A. Teixeira do
(13):1021–1024. crop relatives: Genomic and breeding re- Amaral, Jr., and M.G. Pereira. 2016. Heter-
Endt, D.J.W. 1983. Where the babaco began. sources: Tropical and subtropical fruits. osis and genetic diversity for selection of
Southern Hort. 13:24–26. Springer, Heidelberg, Germany. papaya hybrids for resistance to black spot
Flindt, M.L.H. 1978. Respiratory hazards from Scheldeman, X., P. Van Damme, J. Alvarez, and and phoma spot. Trop. Plant Pathol. 41:380–
papain. Lancet 311(8061):430–432. J. Romero Motoche. 2003. Horticultural 389.

1064 HORTSCIENCE VOL. 52(8) AUGUST 2017

You might also like